Quipuscoa & Dillon: Cuatro nuevas especies endémicas de Nolana (Solanaceae-Nolaneae) de Arequipa, Perú

Arnaldoa 25 (2): 295-322, 2018 ISSN: 1815-8242 (edición impresa)

[Link] ISSN: 2413-3299 (edición online)

Four new endemic species of Nolana

(Solanaceae-Nolaneae) from Arequipa, Peru

Cuatro nuevas especies endémicas de Nolana

(Solanaceae-Nolaneae) de Arequipa, Perú

Victor Quipuscoa Silvestre

Departamento de Biología, Universidad Nacional de San Agustín,
Arequipa, PERÚ
vquipuscoa@[Link]

Michael O. Dillon
The Field Museum, Integrative Research Center
1400 South Lake Shore Drive, Chicago, IL 60605-2496, USA
mdillon@[Link]

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Este es un artículo de acceso abierto bajo la licencia CC BY-NC 4.0: [Link]
 Quipuscoa & Dillon: Cuatro nuevas especies endémicas de Nolana (Solanaceae-Nolaneae) de Arequipa, Perú

Recibido: 10-III-2018; aceptado: 20-IV-2018; publicado online: 30 –VII-2018; publicado impreso: 30-VIII-2018
Abstract
In preparation of a monographic treatment for Nolana L. ex L. f. (Solanaceae-Nolaneae), four
new species are described from department of Arequipa, southern Peru: N. bombonensis Quip. &
M. O. Dillon, prov. Islay, district of Punta de Bombón, Lomas de Alto La Punta; N. callae Quip. &
M. O. Dillon, prov. Islay, district of Punta de Bombón, Lomas de Jesús; N. quicachaensis Quip. & M.
O. Dillon, prov. Caravelí, dist. Quicacha; and N. tricotiflora Quip. & M. O. Dillon, prov. Camaná,
dist. Quilca, Lomas de Quilca. These species are diagnosed, described, illustrated and compared
to nearest geographic neighbors in southern Peru. To aid in recognition, a key to Nolana species
reported from Arequipa is provided.

Keywords: Nolana, Nolaneae, new species, department of Arequipa, Peru, Solanaceae.

Resumen
En la preparación para la publicación de la monografía de Nolana L. ex L. f. (Solanaceae-
Nolaneae), se describen cuatro especies nuevas en departamento de Arequipa del sur de Perú: N.
bombonensis Quip. & M. O. Dillon, prov. de Islay, distrito de Punta de Bombón, Lomas de Alto La
Punta; N. callae Quip. & M. O. Dillon, prov. de Islay, distrito de Punta de Bombón; N. quicachaensis
Quip. & M. O. Dillon de la prov. Caravelí, dist. Quicacha; y N. tricotiflora Quip. & M. O. Dillon
de la prov. Camaná, dist. Quilca, Lomas de Quilca. Además de las diagnosis y descripciones, se
realizan las ilustraciones y se comparan con las especies geográficamente vecinas más cercanas del
sur de Perú. Para ayudar al reconocimiento, se proporciona una clave para las especies de Nolana
reportadas en departamento de Arequipa.

Palabras clave: Nolana, Nolaneae, endémicas, especies nuevas, departamento de Arequipa, Perú,
Solanaceae.

Citación: Quipuscoa, V. & M. O. Dillon. 2018. Four new endemic species of Nolana (Solanaceae-
Nolaneae) from Arequipa, Peru. Arnaldoa 25 (2): 295-322. doi: [Link]
arnaldoa.252.25201

Introduction 50 kms of the Pacific Ocean (Rundel et al.,

1991; Dillon, 1997; Dillon et al., 2003). Only
Nolana L. ex L. f. (Solanaceae-Nolaneae)
a few species are distributed above 2000 m
is a genus consisting of 89 species,
and/or at a distances of 50-500 kms inland
including the four described here (Dillon,
from the coast, e.g., N. chapiensis M. O.
2016, Table 1). No fewer than 39 species
Dillon & Quip., N. confinis I. M. Johnst., N.
have been reported from Peru, and another
laxa (Miers) I. M. Johnst., N. lezamae M. O.
three species which have distributions
Dillon, Leiva, & Quip., N. urubambae Vargas,
ranging into northern Chile (Nolana
and N. weberbaueri I. M. Johnst.). Most
adansonii (Roem. & Schult.) I. M. Johnst., N.
species are narrow endemics, with small,
gracillima (I. M. Johnst.) I. M. Johnst. and N.
restricted geographic ranges and specific
lycioides I. M. Johnst.). This brings the total
ecological requirements, but a few species
number of Nolana potentially encountered
have larger geographic distributions and
in Peru to 42 species. Of this number,
occur over wide geographic ranges, e.g.,
32 species (75%) are recorded from the
Nolana humifusa (Gouan) I. M. Johnst. and
Department of Arequipa. In Peru, greatest
N. gayana (Gaudich.) Koch.
species diversity is confined to near-
ocean localities termed lomas formations, The area occupied by Nolana species
between 50-800 m elevation and within in the department of Arequipa is an arid

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strip bordering the Pacific Ocean, ca. 500 in easily observed characters and the
kms long and ca. 50 kms wide or an area dramatic loss of discriminant characters
of approximately 25,000 km2 (Dillon, 1997; only observable in living material.
Dillon et al., 2003). The desert is essentially Upon drying, Nolana specimens lose
continuous, but there are changes characters, i.e., working with herbarium
in physiognomy and discontinuities or dried material is much more difficult.
provided by intervening river valleys. In this study and in the preparation of a
The major rivers to dissect the coast are monograph, virtually all taxa have been
Río Ocoña, Río Camaná, Río Quilca, Río examined and photographed in the living
Tambo, and Río Osmore. The distribution state. Recent discoveries during field
of the 32 species recorded from the studies in the Department of Arequipa
department contains some taxa with have led to the recognition of four new
wide distributions, essentially occurring species considered morphologically
throughout, for example Nolana spathulata distinct and geographically circumscribed.
Ruiz & Pav., or the aforementioned species
As illustrated in Dillon et al. (2009),
recorded from northern Chilean localities
members from three clades are represented
(i.e., N. adansonii, N. gracillima, N. lycioides),
in Arequipa, i.e., Clades D, E, and F. It is
but most are narrow endemics.
surmised that the four species described
The amazing species diversity in Nolana here would all fall within Clade F. Clade
has been stimulated by ecological changes F was recovered as a well-supported but
within the study area. Both long-term poorly resolved group (Dillon et al., 2009);
(glacial cycles, ~15,000 years) and short- 27 species confined to Peru, and one
term (ENSO events, ~15 years) causational Chilean species, N. intonsa. Of the new
phenomena (Dillon et al., 2003). Current- species proposed here, only one has been
day distributions belie the dynamic included in phylogenetic analyses (Dillon
history of the coastal region over the last et al., 2007; 2009; Tu et al., 2008). Comments
4 my (Dillon et al., 2009). It is assumed that concerning phylogenetic relationships
taxa are products of allopatric speciation are largely postulated from comparative
models, where isolation plays an important morphology.
role in insuring geographic fidelity in
breeding populations. Today, instances of
sympatry at a specific localities are deamed Materials and methods
to be the result of transport of mericarps
Descriptions were made from both
downslope by rain and wind. While rare,
living material encountered during
the periodic coastal rains move mericarps
field studies and dried herbarium
and provide germination opportunities.
specimens deposited in HSP, F, and
Nolana is easy to identify as a genus USM. All acronyms follow those in
with its unique mericarp fruits (Knapp Index Herbariorum ([Link]
2002); however, species delimitations are [Link]/science/ih/). Conservation
open to interpretation and recognition of status was assigned using IUCN criteria
the number of accepted species has varied (2017) combined with field observations
widely (Mesa, 1981; Dillon et al., 2009). and geographic distribution based on
This variation in the number of species herbarium specimens.
recognized is due to widespread homology

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We utilize the “morphological cluster” apically rounded, the bases cuneate,

concept in recognition of species in Nolana margins revolute, abaxially canaliculate.
(see Mallet, 1995), defined as “assemblages Inflorescences of solitary flowers in upper
of individuals with morphological features leaf axils; pedicels cylindrical, densely
in common and separate from other pubescent, 2-7 (-10) mm long. Flowers
assemblages by correlated morphological 5-merous; calyx narrowly campanulate, 3-4
discontinuities in a number of features”. mm wide at anthesis, densely lanuginous,
In addition to the diagnoses provided for 5-lobed, the tube ca. 3 mm long, 3-5 mm
the new species, specific characters useful in diameter, the lobes oblong-lanceolate,
in recognition of species are detailed in the unequal, 4-5 (-5.5) mm long, 1.5-2 mm
Key to Species of Arequipa. wide, the apices obtuse or rounded;
corollas zygomorphic, infundibuliform,
Resultads and discussion 15-20 mm wide at anthesis, 15-20 (-22) mm
Taxonomic treatment long, light lavender or lilac, the throat clear,
externally and internally glabrous, the
1. Nolana bombonensis Quip. & M.
trichomes uniseriate; stamens 5, included,
O. Dillon, sp. nov. (Fig. 1, 2, 3)
the filaments inserted on lower third of
TYPE: PERU. Arequipa: Prov. Islay, corolla, unequal, 8-14 mm long, pilose at
dist. Punta de Bombón, Lomas de Alto La the bases; anthers dithecal, purple, the
Punta, 17º09´37.0” S, 71º46´35.5’ W, 168 m, thecae ca. 1.2 mm long, ca. 1 mm wide,
31-X-2017, V. Quipuscoa S., M. O. Dillon, glabrous; ovary glabrous, ca. 1 mm long,
C. Tejada P., M. Balvin A., S. Huamaní Q., ca. 1.2-1.5 mm wide, basal nectary ca. 1 mm
M. Bedoya C., C. Sanz N., & M. Flores M. wide, the carpels 5, the style included, (4-)
6338 (holotype: HSP-007821; isotypes: 7-11 mm long, the stigma lateral, green, ca.
F-2322991, USM-301275). 0.5 mm long. Fruits mericarps, 5, 1-seriate,
polyhedrons, brown to black, rugose, 2-2.5
Diagnosis
mm, 3 large, 2 small, included within the
Nolana bombonensis is most similar to expanding calyx, 3-4-seeds.
N. volcanica and differs from that species in
Phenology
cinereous habit, oblong densely lanuginous
leaves, calyx lobes unequal, apically obtuse or Flowering august-november.
blunt, and pale lilac or light lavender corollas
Etymology
lacking prominent dark-purple nectar guides
in the inner throat. The specific epithet is derived from
the geographic area of Punta de Bombón,
Description
near the town of Cocachacra in southern
Sprawling, cinereous, suffrutescent Department of Arequipa.
subshrubs, to 1.5 m in diameter, 25-50 cm
Distribution and ecology
tall; stems intricately branched, prostrate
to decumbent or spreading, stems to Nolana bombonensis has been recorded
50 cm long, much-branched, densely from several locations south of Punta de
lanuginous to arachnoid. Leaves alternate, Bombón, Department of Arequipa (Figure
sessile, the blades linear-oblong, 5-8 (-11) 4) at the mouth of the Río Tambo. The
mm long, 1-2 (-2.5) mm wide, densely type locality is the most northwestern
lanuginous pubescent, succulent, entire, population with additional localities

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extending about 30 kms to the south along N. sedifolia Poepp., and N. villosa (Phil.)
a low strip of land near the ocean. I. M. Johnst.; however, these species
are very different in their floral and
Putative relationships
vegetative morphology; they have no clear
Nolana bombonensis is distinctive among relationships with any Peruvian species
its congeners in Peru with its dense, gray, (Dillon et al., 2009).
tomentose pubescence and lite lavender
Conservation status
corollas. It is apparently a narrow endemic
restricted to a small environmentally Critically Endangered (CR); overall
distinct habitat, and sympatric at some distribution <10 km2 (CR) and perhaps
localities with other Nolana species, <250 individuals. See IUCN (2017) for
e.g., N. adansonii, N. pilosa, N. spathulata, explanation of measurements. Agriculture
and N. thinophila I. M. Johnst. When this and poultry farming is expanding rapidly
plant was first encountered in 2003, it in this area severely impacting coastal
was mistaken for N. volcanica, a species ecosystems; the future of this and other
originally described from above Mollendo, plants is very uncertain.
i.e., Lomas of Yuta (Quipuscoa et al., 2016).
Additional specimens examined
When detailed sampling more clearly
defined the range of phenotypic variation PERU. Arequipa: prov. Islay, Punta de
in N. volcanica, the population at Punta de Bombón. Carretera Costanera, Aprox. 10
Bombón was deemed distinct. Km al Sur de Punta de Bombón, 17º11’S,
71º43’W, 20 m, 19-XI-2005, M. O. Dillon, J.
This species was included in the
Wen, S. Leiva G., V. Quipuscoa S., E. Ortiz,
molecular studies under the name, N.
M. Zapata C., M. Corrales M. & J. Castillo
volcanica (Quipuscoa et al. 2930) and its
8989 (HSP, HUSA, F-2276597), 8995 (HSP,
relationships were with other southern
HUSA, F-2276603); ca. Km 153 carretera
Peruvian species. Utilizing a variety of
Punta de Bombón-Ilo, 17º11´57.6” S,
DNA markers, N. volcanica was recovered
71º42´08.6’ W, 7 m, 28-X-2017, V. Quipuscoa,
with congeners, i.e., GBSSI sequences
M. O. Dillon, M. Balvin A., S. Huamaní Q. &
(Dillon et al., 2007) found N. lycioides as
M. Bedoya C. 6226 (HSP).
its sister taxon; LEAFY second intron (Tu
et al., 2008) recovers it in a clade with N. Notes
cerrateana, N. intonsa (Chilean), and N. Nolana bombonensis was initially
lycioides; and a variety of chloroplast confused with Nolana volcanica Ferreyra
markers recovered it in an unresolved (1960), a species based upon a collection
clade with other over a dozen other species by Ms. Dora B. Stafford (holotype:
from Arequipa (Dillon et al., 2009). K000532281) from a locality ca. 40 kms
The growth form is not unique among north of the Río Tambo. That collection
Nolana, but the gray The character of dense was gathered from the quebrada above
tomentose pubescence is not common in Mollendo at ca. 600 m (2000 ft) from habitats
the genus. Among Peruvian taxa, only of “sand and volcanic ash” in the Lomas
Nolana tomentella Ferreyra shares the of Yuta. Sampling N. volcanica throughout
character. There are tomentose taxa found its range and over a period of years
in Chile, e.g., Nolana diffusa I. M. Johnst., illustrated that the density of pubescence
N. tocopillensis (I. M. Johnst.) I. M. Johnst. is variable with glabrescence typical. The

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floral morphology and corolla coloration apically acute, basally attenuate, slightly
pattern in Nolana volcanica is significantly revolute. Inflorescences of solitary flowers
different from N. bombonensis. In contrast in upper leaf axles; pedicels 5-30 (-35)
to N. bombonensis, N. volcanica is composed mm long, lanuginous. Flowers 5-merous;
of spreading perennials appearing green, calyx campanulate 4-7 (-8) mm wide,
and flowers with attenuate calyx lobes, densely lanuginous, 5-lobed, the tube
and shorter, pale blue corollas with a dark 1.5-2 mm long, 3-5 mm in diameter, the
purple band and nectar guides within the lobes narrowly deltoid to linear, unequal,
throat. 5-10 (-12) mm long, 1-2.5 (-3) mm wide,
the apices acute to obtuse; corollas
2. Nolana callae Quip. & M. O. Dil-
zygomorphic, infundibuliform, 15-23 mm
lon, sp. nov. (Fig. 5, 6)
wide at anthesis, 17-23 (-25) mm long,
TYPE: PERU. Arequipa: Prov. Islay, lavender, inner throat purple, externally
dist. Punta de Bombón, Lomas de Jesús, and internally glabrous; stamens 5,
ca. Km 172 carretera costanera, entre Corío included, the filaments 11-15 (-17) mm
y Yerba Buena, 17º14´39.03” S, 71º32´46.5’ long, 2 long, 3 short, bases pilose; anthers
W, 265 m, 06-I-2018, V. Quipuscoa S., M O. dithecal, purple, the thecae 2-2.5 mm long,
Dillon & C. Tejada P. 6857 (holotype, HSP- ca. 1 mm wide, glabrous; ovary glabrous,
007823; isotypes, F-2322992, USM-301277). ca. 1 mm long, 1.2-1.5 mm wide; nectary
Diagnosis basal, orange, ca. 1 mm wide, the carpels
5, the styles included, 6-9 mm long, the
Nolana callae most closely resembles N. stigma lateral, green, ca. 0.5 mm long.
cerrateana, but differs in characters of the Fruits mericarps, 5, 1-seriate, polyhedrons,
habit, leaves, pedicels, calyx shape, and number black, lightly rugose to smooth, 4-3 mm
of mericarps. The former species has a prostrate long, 2-3 mm in diameter, equal, included
habit forming mats to 1.2 m in diameter; within maturing calyx; 2-3-seeds per
slightly larger elliptic-lanceolate leaves 15- mericarp.
30 mm long, 5-12 mm wide, base of petioles
with conspicuous sheath, abaxial and adaxial Phenology
surfaces lanuginous; pedicles to 35 mm long; Flowering october-january.
calyx with narrowly deltoid to linear lobes and
Etymology
five mericarps.
The specific epithet is dedicated in
Description
homage to the professor of Botany of the
Suffruticose perennials, 0.7-1.2 m in National University of San Agustin de
diameter; stems decumbent to repent, Arequipa, Abraham Calla Paredes, for his
much-branched, lanuginose, glabrescent, dedication to the teaching of algae and
young branches greenish-purple, shared friendship for many years.
pubescence of simple trichomes. Leaves
Distribution and ecology
alternate, petiolate, swollen bases 2-4
mm wide, sclerified; petioles slightly Nolana callae is considered endemic to
ribbed, 3-7 (-10) mm long, lanuginous, Arequipa and is restricted to dry, rocky
the blades elliptic to lanceolate, 15-30 slopes at the lower part of the Lomas de
(-40) mm long, 5-12 (-15) mm wide, laxly Jesus, between Punta de Bombón and
lanuginous, glabrescent, succulent, entire, Ilo (Figure 4). To date, it has only been

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recorded from the type locality together prominent dark band and nectar guides in
with Nolana adansonii, N. bombonensis, N. the throat of the corolla (Figure 6B).
spathulata, and Solanum peruvianum L. It was
3. Nolana quicachaensis Quip. &
discovered in disturbed roadside localities
M. O. Dillon sp. nov. (Fig. 7, 8)
and likely, with continued exploration, it
is anticipated that the distribution may be TYPE: PERU. Arequipa: Prov. Caravelí;
expanded upslope. Quicacha, entre Caramba y Quicacha,
15º39´24.58” S, 73º48´53.07’ W, 1593 m, 30
Putative relationships
-XI-2017, V. Quipuscoa S., M. O. Dillon, M.
Nolana callae has not been included Balvin A., S. Huamaní Q, M. Bedoya C. & W.
in phylogenetic analysis and its putative Ancalla Ch. 6763 (holotype, HSP-007825;
relationships are here based upon isotypes, F-2322993, USM-301276).
comparative morphology and distribution.
Diagnosis
It has similarity with N. cerrateana, sharing
habit and lanuginous leaves; however, N. Nolana quicachaensis most closely
cerrateana has longer pedicels to 50 mm, resembles N. lycioides, with its woody, much-
more fasciculate leaves, and a calyx with branched habit and tubular corollas; however,
purple coloration, and 10-14 mericarps. the new species has fasciculate, sessile, terete
leaves, pubescent with short glandular
Conservation status
trichomes; solitary axillary flowers forming a
Critically Endangered (CR); overall weak, terminal raceme; calyx lobes narrowly
distribution <10 km2 (CR) and perhaps deltoid, long-attenuate, 4-6 mm long; the
<250 individuals. See IUCN (2017) for corollas hypocrateriform with yellow tube and
explanation of measurements. white lobes; 15-18 mericarps.
Additional specimens examined Description
PERU. Arequipa: Prov. Islay, Dist. Shrubs, 0.3-1 m tall, the stems of dense
Punta de Bombón. Punta de Bombón, wood, 3-4 cm in diameter, lenticellate,
17º14.6’S, 71º32.7’W, 27-X-2003, V. fissured, much-branched, glabrescent.
Quipuscoa S., M. O. Dillon, R. Freyre, & Leaves fasciculate, sessile, the blades linear
M. Benavides 2930 (HSP, HUSA), 2933 to narrowly spathulate, 5-9 (-11) mm long,
(HSP, HUSA); Prov. Islay, Dist. Punta 0.5-1.5 (-2) mm wide, sigmoid, glutinous,
de Bombón, Lomas de Jesús, 17º14´40.7” succulent, entire, apically cuspidate,
S, 71º32´46.06’ W, 267 m, 29-X-2017, V. basally attenuate, terete. Inflorescences
Quipuscoa S., M.O. Dillon, C. Tejada P., M. weakly racemose, the flowers solitary,
Balvin A., S. Huamaní Q. & M. Bedoya C. axillary, pedicles cylindrical, glandular-
6309 (HSP, F). pubescent, 5-8 (-13) mm long. Flowers
Notes 5-merous; calyx narrowly campanulate,
3-4 mm wide, glutinous inside and out,
Nolana callae most closely resembles 5-lobed, the tube ca. 1 mm long, 2.5-3 mm
N. cerrateana, a species from the area in diameter, the lobes linear-lancelate,
of Camaná, further north in Arequipa; unequal, 4-5 (-6) mm long, 1-1.5 mm
however, it also shares some superficial wide, apices long-attenuate; corollas
similarity to N. intonsa I. M. Johnst. from zygomorphic, tubular-hypocrateriform,
northern Chile. These species also have a 10-12 mm wide at anthesis, 18-20 (-23)

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mm long, the tube yellow, the lobes white, comparative morphology and distribution.
externally glutinous internally glabrous; It has similarity with N. lycioides, but differs
stamens 5, included, the filaments inserted in a range of characters.
in the middle of the corolla, unequal, 12-
Conservation status
16 (-18) mm long, 2 long and 3 short, bases
glabrescent; anthers dithecal, white, the Critically Endangered (CR); overall
thecae 1.5-2.5 mm long, ca. 1 mm wide, distribution <10 km2 (CR) and perhaps
glabrous; ovary glabrous, ca. 1 mm long, <250 individuals. Before rational status
1-1.5 mm wide, nectary basal, orange, ca. can be determined, further studies in the
1 mm wide, the carpels 15-18, the styles area are needed to determine population
14-17 mm de largo, the stigma lateral, size and distribution.
ca. 0.5 mm long. Fruits mericarps, 15-18, 4. Nolana tricotiflora Quip. & M.
2-3-seriate, 5 large, 2-2.5 mm long, 1.8-2 O. Dillon. sp. nov. (Fig. 9, 10)
mm in diameter; 10 intermediate, 1.5-1.8
mm long, ca. 1 mm in diameter, 1-3 small, TYPE: PERU. Arequipa: Prov. Camaná,
1-1.2 mm long, 0.5-0.8 mm in diameter, Quilca, Lomas de Quilca, Km 62 carretera
pyriform, oblong, black, lightly rugose Costanera, Carrizales, 16º51´18.6” S,
to smooth, contained within the calyx at 72º12´21.4’ W, 622 m, 03-XI-2017, V.
maturity, 1 seed per mericarp. Quipuscoa S., M. O. Dillon, M. Balvin A., S.
Huamaní Q. & M. Bedoya C, 6433 (holotype,
Phenology HSP-007827; isotypes, F2322994, USM-
Flowering october-december. 301274). Figure 9

Etymology Diagnosis

The specific epithet is derived from Nolana tricotiflora differs from all other
the geographic area of Quicacha, near the members of the genus with a unique combination
town of Cháparra in north Department of of characters not encountered. Its erect crooked,
Arequipa. woody trunks to 50 cm tall; numerous spirally-
arranged leaves, and terminal, three-branched,
Distribution and ecology
scorpioid cymes.
Nolana quicachaensis is only known
Description
from the type between the towns of
Caramba and Quicacha (Figure 4). It was Shrublets, 30-50 cm tall; stems woody,
found growing between granitic rocks 1-2 cm in diameter, brown, lenticellate,
in the lower part of the south-facing cracked, slightly branched at the base,
slopes. Associates included members pubescent on mature stems, densely villous,
of desert vegetation such as, species of young stems greenish-purple, pubescent
Cactaceae (Melocactus, Cumulopuntia, with crooked glandular trichomes to 2 mm
Weberbauerocereus), Asteraceae (Helogyne, long. Leaves whorled, crowded at branch
Baccharis) and annual grasses. apex, sessile to subsessile, the leaf blades
elliptic to linear-lanceolate, 10-14 (-15) mm
Putative relationships
long, 2-3 (-3.5) mm wide, falcate, distally
Nolana quicachaensis has not been flexed upwards, both surfaces villous, the
included in phylogenetic analysis and its trichomes to 1.5 mm long, apically capitate-
putative relationships are here based upon glandular, succulent, entire, apically acute

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to obtuse, basally attenuate. Inflorescences on ocean-facing slopes in the Carrizales

of 3 terminal, lax scorpioid cymes, 7-12 sector. Vegetative associates include
cm long, each with ca. 12 flowers; pedicels Heliotropium (Boraginaceae), Cryptantha
villous, apically capitate-glandular, 2-3(-4) (Boraginaceae), Palaua (Malvaceae),
mm long, subtended by reduced, villous, Spergularia (Caryophyllaceae), Senecio
foliar bracts. Flowers 5-merous; calyx (Asteraceae), Oenothera (Onagraceae), Nasa
campanulate, 8-12 mm wide at anthesis, (Loasaceae), Hoffmannseggia (Fabaceae),
externally villous, glutinous, internally Pasithea, Cytharexylum (Verbenaceae),
pubescent, 5-lobed, the tube 3-4 mm long, Hierobotana (Verbenaceae), Loxanthocereus
3-5 mm in diameter, the lobes deltoid, (Cactaceae) and Vasconcellea (Caricaceae).
unequal, 7-10 (-12) mm long, 3-4 mm
wide, apically acute to obtuse; corollas Putative relationships
zygomorphic, infundibuliform, 17-22 (-25) Nolana tricotiflora has not been included
mm wide at maturity, 17-20 (-25) mm in phylogenetic analysis and its putative
long, purple, the throat with dark nectar relationships are difficult to establish upon
guides, externally and internally glabrous; comparative morphology and distribution.
stamens 5, included, the filaments 9-12
Conservation status
(-13) mm long, 2 long, 3 short, bases pilose;
anthers dithecal, purple, the thecae 1.5-2.5 Critically Endangered (CR); overall
mm long, ca. 1 mm wide, glabrous; ovary distribution <10 km2 (CR) and perhaps
glabrous, ca. 1.5 mm long, 1.5-2 mm wide, <250 individuals. See IUCN (2017) for
nectary basal, orange, ca. 1 mm wide, the explanation of measurements.
carpels (11-) 12-18, the styles included,
Notes
flat, 9-12 mm long, ca. 1 mm wide, the
stigma apical, ca. 0.5 mm long, 1-1.5 mm Nolana tricotiflora contains a combination
wide. Fruits mericarps, 12-18, 2-seriate, of characters not to be met in any other
pyriform, black, 5 large, 2-2.5 mm long, member of the genus. No woody species
1.5-2 mm in diameter; 10 medium, 1-3 approach its overall habit with crowded
small, lightly rugose to smooth, included cauline leaves and very long villous
within the expanded calyx as it matures; pubescence with glandular apical cells. But
larger mericarps with 2-3-seeds. most unusual, is the inflorescence of three-
branched weak scorpioid cymes with
Phenology
large flowers. In the majority of Peruvian
Flowering october-november. Nolana species, the flowering stems are
unmodified and flowers are borne as
Etymology
solitary in leaf axils. The only exceptions
The specific epithet refers to the are southern Peruvian species, which have
inflorescence of three terminal, lax modifications of flowering stems into
scorpioid cymes. recognizable inflorescences with modified
Distribution and ecology bracts subtending individual flowers, but
arising from a basal rosette of modified
Nolana tricotiflora has only been recorded leaves, N. inflata and N. weissiana. In N.
from the type locality, the Lomas of Quilca, scaposa, the condition reaches it maximum
between Matarani and Quilca (Figure 4). development where the inflorescence is a
It was found growing between 550-800 m modified branch with subtending floral

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bracts. None of these species remotely resemble N. tricotiflora.

Key to Nolana species recorded from Department of Arequipa

1 Plants with persistent basal rosette of petiolate leaves; 2

leaves
_ Plants without persistent basal rosette of petiolate leaves; 4
if ephemeral central rosette present, then leaves similar
to cauline leaves in size and shape
2 Basal leaves wtih strictly entire blades; flowering shoots N. scaposa
with sessile oval to ovate bracts subtending flowers, the
flowers congested in dense inflorescence, densely villous
pubescence; calyx campanulate, deeply lobed, lobes
unequal, obtuse; corollas white
_ Basal leaves with dentate to crenate blades; flowering 3
shoots with petiolate, lanceolate bracts subtending
flowers, the inflorescence open, flowers not congested,
densely hirsute-glandular to arachnoid or lanuginous;
calyx globose or bladder-inflated, lobes equal, triangular,
acute; corollas deep purple to lavender
3 Cauline leaves ovate N. inflata
_ Cauline leaves laneolate N. weissiana

4 Inflorescences well-developed, consisting of three, N. tricotiflora

terminal, weak scorpioid cymes
_ Inflorescences not present, flowers usually solitary in 5
upper leaf axles, weaking racemose, but never scorpioid
cymes
5 Calyx appearing zygomorphic 6
_ Calyx actinomorphic or simply bilobed, the apices of 8
lobes triangular to attenuate, never expanded apically
6 Leaf blades lancolate N. pallidula
_ Leaf blades broadly lanceolate, spathulate, ovate 7

7 Blades with bases truncate N. spathulata

_ Blades cuneate N. arenicola

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8 Leaves clearly petiolate, the bases auriculate, at times N. adansonii

connate, the blades cordiform, more rarely reniform to
lanceolate or elliptic, glabrous, surface with salt glands
that attract atmospheric moisture causing the look and
feel of oil
_ Leaves not obviously petiolate, the bases not auriculate 9
nor connate, the blades linear, lanceolate, elliptic to ovate,
but never cordiform, surfaces glabrous to glabrescent to
lanuginous or tomentose with stellate pubescence, never
oily
9 Leaves linear, oblong, oblanceolate or narrowly 10
spathulate
_ Leaves laminar, elliptic, lanceolate to ovate, mostly 2.5 21
mm wide or greater, margins often revolute

10 Leaves glabrous 11
- Leaves pubescent with stipitate-glandular trichomes, 16
villous, lanuginous or tomentose
11 Calyx bilobed, cylindrical, 2-4 mm wide, apically with 12
1-2 deep clefts, the proper calyx lobes reduced, either
absent or inconspicuous 2-3 mm wide
- Calyx uniformly 5-lobed, campanulate-tubular to globose 14
12 Minute erect annuals, 2-5 cm tall; flowers blue (Camaná) N. minor
_ Spreading robust, annuals to perennials, generally 20-30 13
cm tall; flowers violet, purple, to white
13 Leaves 10-20 mm long, 2-4 mm wide, obovate to N. arequipensis
oblanceolate, corollas white, 3-6 mm wide, distributed in
inland habitats (100-300 m)
_ Corollas purple to violet or lavender, 10-40 mm wide. N. thinophila
Calyx glabrous to short, stipitate glandular, corolla ca. 1
cm broad (beachside)
14 Calyx globose, 6-10 mm tall, 4-8 mm wide; leaves linear N. chancoana
to oblanceolate, 10-40 mm long, 2-6.5 mm wide
- Calyx cylindrical-tubular to campanulate, 4-9 mm tall, 15
1.5-3.5 wide; leaves 8-25 mm long, 0.8-1.5 mm wide
15 Leaves lanuginous and glabrescent; flowers sessile or N .
with short pedicels; mericarps 2 spergularioides

_ Leaves strictly glabrous; flowers pedicles, 5-13 mm long; N. gracillima

mericarps 5

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16 Stems, leaves and calyx lobes tomentose, canescent or N. bombonensis

gray; leaves linear-oblong, mm long, mm wide

_ Stems, leaves and calyx lobes stipitate-glandular or 17

lanuginous, green; leaves elliptic to ovate

17 Pubescence of stipitate-glandular trichomes 18

_ Pubescence lanuginous 20

18 Perennial herbs; corolla 10-15 long, 4.8-9 mm wide; N. confinis

leaves 6-14 mm long, 1.6-1.8 mm wide
- Shrubs, much-branched; corollas infundibularis or 19
salverform, generally longer than 15 mm
19 Corollas infundibular, 13-27 mm long, purple N. lycioides
- Corollas salverform, 18-23 mm long, white N .
quicachaensis
20 Leaves linear-lanceolate, 7-10 mm long, 1.2-2.8 mm N. volcanica
wide, pubescence lanuginous, sometimes dense
_ Leaves linear, 8-16 mm long, 1-2 mm wide, pubescence N. tovariana
laxly pubescent
21 Leaves pubescent with stellate pubescence N. pallida
- Leaves glabrous or variously pubescent, never stellate 22
22 Leaves glabrous 23
- Leaves pubescence 24
23 Corolla 20-30 mm long; calyx prominently pleated at the N. coronata
sinus, evenly rounded off beneath it, glabrous

- Corolla 15-20 mm long; calyx lobes deeply cut, strigose N latipes

24 Leaves with short or long stipitate-glandular trichomes 25
- Leaves pubescent of non-stipitate trichomes, tomentose 28
to strigose or sericeous
25 Pubescent with short stipitate-glandular trichomes 26
- Pubescence villous, trichomes long, terminally stipitate- 27
glandular
26 Calyx bilabiate, the lobes connate, 2 and 3 fused, 15-26 N. aticoana
mm; corollas 28-50 mm long, 25-60 mm wide; mericarps
3-5

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- Calyx zygomorphic with all lobes on one side, 18-30 N. mariarosae

mm long; corollas 32-38 mm long, 24-30 mm wide;
mericarps 15-19
27 Leaves elliptic, 8-15 mm long, 2-3.5 mm wide N. chapiensis

- Densely pubescent with long, stipitate-glandular, N. pilosa

trichomes; calyx flat, corolla 13-22 mm broad; leaves 10-
27 mm long, 1-6 (-8) mm wide; stems and leaves pilose
28 Pubescence stiff 29
- Pubescence tomentose to lanuginous 30
29 Leaves Leaves lanceolate to oblanceolate or spathulate, N. johnstonii
15-30 mm long, 10-20 cm wide, strigose
- Leaves lanceolate to narrowly ovate, 10-65 mm long, 13 N. plicata
mm wide; trichomes stiff (shaggy) erect sericeous

30 Leaves canescent, pubescence tomentose, trichomes N. tomentella
inter-tangled and arachnoid
- Leaves appearing green, pubescence lanuginous, the 31
hairs not inter-tangled, never arachnoid
31 Leaves 15-30(-40) mm, 5-12 (-15) mm wide; corollas N. callae
15-23 mm wide; mericarps 5
- Leaves 11-26 mm long, 2-5.5 mm wide; corolla 12-20 N. cerrateana
mm wide; mericarps 10-14

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Acknowledgments Literature cited

Curators and collection managers Dillon, M. O. 1997. Lomas Formations-Peru. Pp 519–
at HUSA, HSP, HUT, USM are thanked 527. In: Davis SD, Heywood VH, Herrera-McBryde
O, Villa-Lobos J, & Hamilton AC (Eds), Centres of
for providing loans and permitting Plant Diversity, A Guide and Strategy for their Con-
examination of collections. Field studies servation. WWF, Information Press, Oxford.
were supported, in part, by grants to Dillon, M. O. 2005. Solanaceae of the Lomas forma-
VQS from CIENCIACTIVA-UNSA tions of Coastal Peru and Chile. Pp 131–155. In:
investigation project No 37-2016-UNSA. Hollowell V, Keating T, Lewis W, & Croat T (Eds),
VQS thanks the Department of Biology, A Festschrift for William G. D’Arcy: The Legacy of
a Taxonomist. Mono. Syst. Bot. Ann. Missouri Bot.
UNSA for the permits granted to carry out
Gard. 104.
continuous field studies and the members
Dillon, M. O. 2016. 71. Nolana (Solanaceae). Pp
of IMOD (Michael Owen Dillon Institute)
343–344. In: Barboza GE., Hunziker AT, Bernarde-
to the entire Project team of endemic llo G, Cocucci AA, Moscone AE, Carrizo Garcia C,
species: Felipe Sinca, Károl Durand, Fuentes V, Dillon MO, Bittrich V, Cosa MT, Subils R,
Daniel Ramos, Rafael Pérez, Italo Treviño, Romanutti A, Arroyo A, & Anton A, The Families
Margarita Balvín, Susan Huamaní, and Genera of Vascular Plants. Asterales. Vol. 8.
Springer Verlag, Berlin.
Geraldine Rosado, Maricruz Bedoya,
Cristian Tejada, Raquel Medina, Wendy Dillon, M. O.; M. Nakazawa & S. Leiva. 2003. The Lo-
mas Formations of Coastal Peru: Composition and
Ancalla, Claudia Sanz, and Martín Flores, Biogeographic History. Pp 1–9, In: Haas J, Dillon
for their collaboration in the conception of M. O (Eds), “El Niño in Peru: Biology and Culture
the project, obtaining data in the field and Over 10,000 Years.” Fieldiana: Botany, N. S. 43,
work in the herbarium. publ. 1524.
Dillon, M. O.; T. Tu; A. Soejima; T. Yi; Z. Nie; A. Tye
Contribution of the authors
& J. Wen. 2007. Phylogeny of Nolana (Nolaneae,
V.Q.: Principal investigator, co- Solanoideae, Solanaceae) as inferred from granule-
bound starch synthase I (GBSSI) sequences. Taxon
ordinator of the fieldwork in the data
54: 1000–1011. DOI: 10.2307/25065900 http://
collection for the taxa, ecological data, [Link]/stable/25065900
photography and botanical collection;
Dillon, M. O.; T. Tu; L. Xie; S. Quipuscoa & J. Wen.
responsible of the descriptions and the 2009. Biogeographic diversification in Nolana (So-
first script, making corrections until the lanaceae), a ubiquitous member of the Atacama
final version. M.O.D.: Co-investigator, and Peruvian Deserts along the western coast of
South America. Journal of Systematics & Evolu-
contributed in the data collection at the
tion, 47: 457–476. [Link]
field trips for describing the taxa, redaction, 6831.2009.00040.x
geographical data collection, photography,
Ferreyra, R. 1955. Nuevas especies de Nolana del
botanical collection, making of the figures, Perú. Publicaciones Museo Historia Natural “Javier
taxonomic key and manuscript correction. Prado” 10: 1–15.
Conflict of interests Ferreyra, R. 1961. Revisión de las especies peruanas
del género Nolana. Memorias Museo Historia Natu-
The authors declare not to have ral “Javier Prado” 12: 1–53.
conflicts of interests.
Ferreyra, R. 1974. Una nueva especie de Nolana para
el Perú. Boletín Sociedad Peruana Botánica 7(1-2):
3–5.
IUCN. 2017. The IUCN Red List of Threatened Species.
Version 2017-3. <[Link]

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tic/categories_criteria_3_1> Downloaded on 02
March 2018.
Johnston, I. M. 1936. A study of the Nolanaceae. Con-
tributions Gray Herbarium 112: 1–83.
Knapp, S. 2002. Tobacco to tomatoes: a phylogenetic
perspective on fruit diversity in the Solanaceae.
Journal Experimental Botany 53: 2001–2022.
[Link]
Mallet, J. 1995. A species definition for the modern
synthesis. Trends in Ecology and Evolution 10:
294–299. doi: 10.1016/0169-5347(95)90031-4.
Mesa, M. A. 1981. “Nolanaceae.” Flora Neotropica 26:
1–197. [Link]
Quipuscoa, S. V.; P. C. Tejada; A. C. Fernández; V. K.
Durand; T. A. Pauca & M. O. Dillon. 2016. Diver-
sidad de plantas vasculares de las Lomas de Yuta,
Provincia de Islay, Arequipa, Perú / Diversity of
vascular plants in Lomas de Yuta, Islay province,
Arequipa, Peru. Arnaldoa 23(2): 517–546.
Rundel, P. W.; M. O. Dillon; B. Palma; A. H. Mooney;
S. L. Gulmon & J. R. Ehleringer. 1991. The phyto-
geography and ecology of the coastal Atacama and
Peruvian Deserts. Aliso 13: 1--50. doi:10.5642/
aliso.19911301.02
Tu, T.; M. O. Dillon; H. Sun & J. Wen. 2008. Phylo-
geny of Nolana (Solanaceae) of the Atacama and
Peruvian Deserts inferred from sequences of four
chloroplast markers and the nuclear LEAFY second
intron. Molecular Biology Evolution 49: 561–573.
doi:10.1016/[Link].2008.07.018

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Table 1. Alphabetical List of Accepted Names and Authorities, Distribution, and

phylogenetic position as suggested by membership in clades of Nolana in South America.
Membership in clades is adapted from Dillon et al 2019. [* designates distribution recorded
from Department of Arequipa]
Species Distribution Clade
1 N. acuminata (Miers) Miers ex Dunal Chile B
2 N. adansonii (Roem & Schult.) [Link]. Chile-Peru* F
3 N. aenigma [Link] & Quip. Peru F
4 N. albescens (Phil.) [Link]. Chile G
5 N. aplocaryoides (Gaudich.) [Link]. Chile G
6 N. arenicola [Link]. Peru* D
7 N. arequipensis [Link] & Quip. Peru* F
8 N. aticoana Ferreyra Peru* F
9 N. baccata (Lindl.) Dunal Chile B
10 N. balsamiflua (Gaudich.) Mesa Chile C
11 N. bombonensis Quip. & [Link] Peru* F
12 N. callae Quip. & [Link] Peru* F
13 N. carnosa (Lindl.) Miers ex Dunal Chile C
14 N. cerrateana Ferreyra Peru* F
15 N. chancoana [Link] & Quip. Peru* D
16 N. chapiensis [Link] & Quip. Peru* D
17 N. clivicola ([Link].) [Link]. Chile E
18 N. coelestis (Lindl.) Miers ex Dunal Chile C
19 N. confinis [Link]. Peru* F
20 N. coronata Ruiz & Pav. Peru* F
21 N. crassulifolia Poepp. Chile G
22 N. dianae [Link] Chile G
23 N. diffusa [Link]. Chile G
24 N. divaricata (Lindl.) [Link]. Chile G
25 N. elegans (Phil.) Reiche Chile B
26 N. filifolia (Hook. & Arn.) [Link]. Chile C
27 N. foliosa (Phil.) [Link]. Chile E
28 N. galapagensis (Christoph.) [Link]. Ecuador D
29 N. gayana (Gaudich.) Koch Peru F

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30 N. glauca ([Link].) [Link]. Chile G

31 N. gracillima ([Link].) [Link]. Chile–Peru* E
32 N. humifusa (Gouan) [Link]. Peru F
33 N. incana (Phil.) [Link]. Chile G
34 N. inconspicua ([Link].) [Link]. Chile G
35 N. inflata Ruiz & Pav. Peru* D
36 N. insularis ([Link].) [Link]. Peru D
37 N. intonsa [Link]. Chile F
38 N. jaffuelii [Link]. Chile (no B
modern Peruvian
records)
39 N. johnstonii Ferreyra Peru* F
40 N. lachimbensis [Link] & Luebert Chile G
41 N. latipes [Link]. Peru* D
42 N. laxa (Miers) [Link]. Peru D
43 N. leptophylla (Miers) [Link]. Chile G
44 N. lezamae [Link], [Link] & Quip. Peru F
45 N. linearifolia Phil. Chile G
46 N. lycioides [Link]. Chile-Peru* D
47 N. mariarosa Ferreyra Peru* F
48 N. minor Ferreyra Peru* F
49 N. mollis (Phil.) [Link]. Chile G
50 N. onoana [Link] & Nakazawa Chile G
51 N. pallida [Link]. Peru* F
52 N. pallidula [Link]. Peru* D
53 N. paradoxa Lindl. Chile B
54 N. parviflora (Phil.) Phil. Chile B
55 N. patula (Phil.) Mesa ex [Link] Chile G
56 N. pearcei [Link]. Peru F
57 N. peruviana (Gaudich.) [Link]. Chile G
58 N. philippiana [Link] & Luebert Chile G

59 N. pilosa [Link]. Peru* F

60 N. platyphylla ([Link].) [Link]. Peru D

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61 N. plicata [Link]. Peru* D

62 N. pterocarpa Phil. ex Wettst. Chile B
63 N. quicachaensis Quip. & [Link] Peru* F
64 N. ramosissima [Link]. Chile G
65 N. reichei M.O. Dillon & Arancio Chile B
66 N. rhombifolia Marti. & Quez. Chile D
67 N. rostrata (Lindl.) Miers ex Dunal Chile C
68 N. rupicola Gaudich. Chile B
69 N. salsoloides (Lindl.) I.M. Johnst. Chile G
70 N. scaposa Ferreyra Peru* D
71 N. sedifolia Poepp. Chile G
72 N. sessiliflora Phil. Chile A
73 N. spathulata Ruiz & Pav. Peru* D
74 N. spergularioides Ferreyra Peru* E
75 N. sphaerophylla (Phil.) Mesa ex [Link] Chile G
76 N. stenophylla I.M. Johnst. Chile C
77 N. tarapacana (Phil.) [Link]. Chile E
78 N. thinophila [Link]. Peru* F
79 N. tocopillensis ([Link].) I.M. Johnst. Chile G
80 N. tomentella Ferreyra Peru* F
81 N. tovariana Ferreyra Peru* F
82 N. tricotiflora Quip. & [Link] Peru* F
83 N. urubambae Vargas Peru F
84 N. villosa (Phil.) [Link]. Chile G
85 N. volcanica Ferreyra Peru* F
86 N. weberbaueri [Link]. Peru D
87 N. weissiana Ferreyra Peru* D
88 N. werdermannii [Link]. Chile G
89 N. willeana Ferreyra Peru D

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Fig. 1. Holotype of Nolana bombonensis, Quipuscoa et al. 6338 (HSP-007822).

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Fig. 2. Nolana in the beaches of Punta de Bombón. A1. N. bombonensis and A2. N. pilosa.B.
Individual of N. bombonensis growing in sand near the ocean with Massiel Nataly Corrales
Medina.

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Fig. 3. Nolana bombonensis. A. Flowering branch, scale bar = 5 mm (Dillon et al. 8989, 19
Nov 2005); B. Flowering stem with corolla, scale bar = 8 mm; C. Mericarps, scale bar = 5
mm.

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Fig. 4. Distribution map for southern Peruvian species of Nolana, N. bombonensis (purple
circle), N. callae (red square), N. quicachaensis (blue hexagon), and N. tricotiflora (green
polygon).

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Fig. 5. Holotype of Nolana callae, Quipuscoa et al. 6657 (HSP-007823).

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Fig. 6. Nolana callae A Flowering individual with Cristian Tejeda-Perez; B. Corollas, scale
bar = 5 mm; C. Mericarps, scale bar = 10 mm (Quipuscoa et al. 6857, 6 Jan 2018).

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Fig. 7. Holotype of Nolana quicachaensis, Quipuscoa et al. 6763 (HSP-007825).

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Fig. 8. Nolana quicachaensis. A. Flowering shrub; B. Frontal view of corollas, scale bar = 8
mm; C. Lateral view of apex, scale bar = 8 mm; D. Mericarps, scale bar = 5 mm.

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Fig. 9. Holotype of Nolana tricotiflora, Quipuscoa et al. 6433 (HPS-007827).

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Fig. 10. Nolana tricotiflora. A. Flowering shrub next to MOD; B .Close-up of shrubby habit,
scale bar = 10 cm; C. Close-up of leafy stem, scale bar = 5 cm; D. Lateral view of flower,
scale bar = 10 mm; E. Frontal view of corolla, scale bar = 5 mm.

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